A REVISION OF THE GENUS TRICHOCENTRUM (ORCHIDACEAE

Transcription

A REVISION OF THE GENUS TRICHOCENTRUM (ORCHIDACEAE
Li odleyaoa 10(3): 183-2 10. 1995.
A REVISION OF THE GENUS TRICHOCENTRUM
(ORCHIDACEAE: ONCIDlINAE)'
FRANCO PuPULlN
Via Com:ggio 57. 20 149 Milano. lta ly
ABSTRACT: A lIUIooomic revisiOll of the geous Trichou", ...m is preseoted. The hislory of lhe ge ous is
highlighled and phylogeoelic relaliooshi ps are discussed. Tweoly-Ihree spocies are reeogo i!ed. penaioing lO
seve n oalural groups. Characlers of each group are disc ussed aoo keys lO groups aoo spccies are included.
Two new spocies. T. t Slrt lltmst aod T. ",agnal. are desc ri bed.
THE DISCOVER y of ¡he frrs{ species of Tricltocentrum predates ils formal descriplion by sorne 50
years. It was found by Hipólito Ruiz and José
Pavón during the Spani sh Expedici6n Botánica of
1777- 1788 10 the vice-kingdorn of Peru (now
Peru and C hile). A plme of what would later be
descri bed as Triclwcentrum pu(chrum Poepp. &
Endl.. by Isidro de Gálvez, sccond illustrator of
the ex pedition, is kept at Royal Botanic Garden
of Madrid under Ihe name Orchis pllflctata (Real
Jardín Botánico, IV, 1252). Thi s walercolor well
illustrates Ihe habit of the species and has a fl oral
detail showing the column and the spurred !ip
with pedice\. Though it bears no date, the plate
was most likely completed before MaTCh 1778
when Gálvez left Lima and retumed 10 Spain together with Ruiz and Pavón. Unfor1unately this
plate. as well as most of ¡he drawings intended
for Rui z and Pavón's Flora Pe ruviana et Chilensiso was never published ; on ly Ihree volumes of
the projecled series of twenty were printed between 1798 and 1802 (Cabello Martín et al..
1988).
Eduard Friedrich Poeppig and S¡c phan Ladislaus Endlicher proposed the genus Trichocentrllm
in 1837 based on a fl owering plant Poeppig collected in Peru in 1830 near Pampayacu in northem Huánuco prov ince along the eastem slopes of
Cordillera Central. In 1881 Bentham assigned Ihe
genus 10 subtribc Oncidieae, encompassing only
New World genera . La/er Schlechler (1 926) di-
vided the genera of Bentham's Oncidieae into
twelve di fferent subtribes; Trichocentrul1l was referred 10 subtribe Trichocentreae on account of ils
spurred lip, excavated stigma and the abscnce of
a column foot. In Schlechtcr's posthumous classification this subtribe appears closely re lmed to
Comparettieae, from which it may be easily separaled on the basis of Ihe free lateral sepals and
the lack of any appcndages m the base of the Iip
(Schlechler, 1926). This classification was largely
foll owed by Schweinfurth ( 1959). Dressler and
Dodson ( 1960) restored subtribe Oncidiinae (sen su Bentham), dividing it into three major alliances; Trichocellrrum was placed in the Ollcidillm
complex. Dressler ( 198 1), examining phylogenetic relationshi ps within Oncidiinae. considered the
spur of Trichocentrum primiliYc ror Ihe sublribe,
and stressed Ihe close relationshi ps of Trichocelltrum with the so-cal1ed " rat-tail " Ollcidium scclion Cebolleta and " mule-ear" Oncidiums, 01/cidiul1l section Pluritllbercll[WQ. and proposed
these two sections may be scparated as the genus
Lo¡Jhiaris Raf. Willi ams and Dressler (ined.. cited
in Dressler, 198 1) suggested Ihat Ihe whole Oncidium complex evolved (rom a common Trichocelltrll1n-li ke ancestror. Ihe tabula infrastigmatica
being derived fram a partial fusion of the lip with
Ihe column b.15e, as in TrichocCJltrlllfl.
Chase ( 1986, 1986b) proposcd a new realignment of the oncidioid genera based on a series of
previously overlooked andlor underutilized fl oral
and vegetati ve charactcrs, incl uding chromosome
number and li fe history. In Ihis preliminary
scheme TrichocetltruIII is considered to be an
, Gralilud!: is e,.,leOOC:(,! lO lhe curalOfS al AMES. S R. CR.
R.AS. K. P. RPSC, SEl. USJ. aoo W fOl" lhe loan of phoI:ographs and critical specimens. Special I hank~ are ad:nowledgtd lO J. T. Alwood. C. H. Dodson. R. L. r:>ress1er aOO G.
A. Romero for lhei r useful suggcslioras and crilical oommenlS
lO Ihe maouscript.
183
PUPULl N- A REVIS ION OF TRJCH OCE.NTRUM
anomalous genus. The unique combination of floral fea tures. vegelutive morphology and low chromosome numbers (2n = 24, 28) suggesl an evoIUlionary path independent from other oncidioid
orchids. However. the convex Slructure of Triehoeelllmlll stipe is also represented in Oncidiinae in
OllcidilllJl seclions CebolJelll and Pl lIriluberculata. This shared character, togelher with a low
chromosome count. vegetalive architeclure (i.e.
pseudobu lb reduction). and site preferences (i.e.
eliophyt ism) seems lo su ppon Oressler's hypolhesis that placed Tric/ux:elltrum close 10 Olleü!ium
seclions Cebolleta and P/urituberell/aU/ (Dress ler.
198 1). A recent study by Chase and Palmer
( 1992), using chloroplast DNA (cpONA) restriction sites of the Oncidiinae, effecl ive ly reduces
the gap between Ihe two schemes of phyletic relalionships. As suggested by Dressler ( 1981), Tri·
cI/Ocelllrllm is assigned 10 Ihe Lophiari.~ elade. but
on Ihe basis of its low chromosorne number. il
now occupies a derived posilion within the group.
Based on these findings, Oressler ( 1993) treated
Triel/Oeel/lrum as a subgroup of section Plllritll bere/l/tlta. If Ihi s elade were elevated to the rank
of gcnus. TricllOcentrum would havc priority over
L()p"iari.~ (Dressler. 1993). The genus U JIJhiarü,
however, was recenll y rcsurrecled by Braem
( 1993) applying it only to Ihe species of OIlC:idi11m seclion PI/lrituberell/(jf(f. with the exception
of Onddium splel/didulI/ A. Rich. Bracm ( 1993)
did nOI refer the species of Om:idi/llll seCl. Ce/)()IIefll 10 Lophiaris based on differences in vegetalivc habi l and polli nariu m morphology.
Tricl/Oeelltr/lm. as circumscribcd herein . ineludes sorne 23 species of epiphytic plams fou nd
from southcrn Mexico (O Brazil . Boli via and Peru.
The generic name TricllOcel1trum refers to the
presence of a slender and fl exuous spur (from Ihe
Gree k words tricllOS. hair. and kemroll. spur). a
fealure chamCleriSlic of T. ,mlel/rum and ils elose
relat ives. However. on ly half of the Trichocell trum species currently rccognized fi l th is generic
concept. The other spec ies vary greatly in spur
morphology. prescnling a shon and gibbose, plu ri-Iobu late. saccate, or elavate spur. Differences in
spur sh¡¡pe are consislentl y associated wilh gross
fl ora l and micromorphological traits. An ¡¡lIempt
10 group Ihe di fferenl taxa on the basis of such
unique seis of shared flora l features is supported
by Iheir geographic di slribulion Ihal shows al
(east fou r major patterns (Fig. 3, 7. 10. 12).
184
Evolution of the spur in Subtribe Oncidiinae
In spite of the unique habit il shares wilh mem·
bers of Ihe Lophiaris elade, Trichocenlrum presenls a radically different fl oral morphology and
ils pollinalion is carried out by fra grance-collecl·
ing euglossine bees (van der Pijl and Dodson.
1966), whereas Ihe Lophiaris elade has evolved
deceil pollinalion. Furthermore. Ihe spur of Triclux:entmm, a non· functional neclary involving
on ly labellum tissuc. is unique among Ihe Onci diinae. The papi llose- hi rsute inlernal walls of Ihe
spur in sorne Triehocellfrum species (Pupulin
& Mora· Retana. 1994) is also a unique fCiJIUre in
the subtribe. A labellar spur is common in many
va ndo id ge ne ra, rnainly in Angraeci nae and
Aerangidinae and in thei r Neolropical rclatives
(Call/py/oce ll¡T/lm and D endrOI)/¡y lax). and is
found in members of Cyrtopodiinae (Galemldm).
bul il represenls an exceplion among Ihe Onci·
diinae. if not among Ihe whole tribe Maxillarieae
(sellsu Oressler, 1993). A labellar neclar cavily is
present in /..eoehi/lls. GOlliocltilus and IO//Opsü.
but this cavily is mere ly a receplaele 10 colleel
the neClar secreted by Ihe colu rnn. and il is pcrhaps primiti ve in the rodri guezioid group. Also
Ihe presence of a tubular spur is not uncommon
in the Oncidiinae. and il can be fou nd in l1lany
genera such as Rodriguezia. COIIII)(/r ettill , Dia·
del/ium, and Plec:tropho ra. A somewhal rudimenlary cavi ly is al so prcsent in Ihe di slant genus
Brac/¡tia, strictly related to the Ada. Brassill and
Asptuia complex. In Oncidiinae genera. however.
Ihe spur is mainly formed by Ihe panial fusion of
lip base and later¡¡1 sepals, whi le colu mnar lissue
may sOllletimes produce one or more "horns"
wi lhin the sepals cavi ly. as in Ihe case of COII/·
parellia. Scelocl/i/¡u. Diadell ill lll and other rodriguczioid genera. A somewhal complele and
graduate succession of neclary shapes can be observed in Oncidiinae, from the simple neClar cav·
¡Iy of L eoc:!,illI s, Goniocllillls and IOllOI)sis.
through the tubul ar nectar cavi ly of PfectrolJllOra
to Ihe lubular nec tar spur with lip-column
"horns" of COlllpa,.e11ia. Rodrig llezia. See lochi·
hu. etc. (Fig. 1). The characleri slic spur of T,.ichocelltl"UlI/ , derived of labellum tissue alone. is
unique among striclly Neolropical orchids. Recent
molecular studies (Chase and Palmer, 1992) highlight the differenl origin of the Triell(x:emrw1I
spur. grouping logelher all Ihe "spurred" O nci-
PUPULlN- A REVIS ION OF TH/CHOCEl«HUM
Fill. 1. Longitudinal se<:tion of column. lip and lalcral sepals in sorne ··spurred·· Oneidiinac. A. IJrachlio slllf"r~a. Vcnezuela
(fmm DUllstcrvilk & Ganly. 1979); B. uochilus Irlcuspid/llus. Costa Ric~ (adapted fmm Chasc. 1986b); C. Gon;och;/'<$ /f'O°
eh;/;""s, Costa Rica (F. I'"/mll,, /71); D. Rrx/r;g"f'ÚCl rompCK/II. Costu Rica (F. "r'p"lí" s.n.): t:. Sa/ochilus pumgrl/liluis.
Venczucla (adaptcd fmm Dunstervi lle & Caray. 1979); t ComfN,rmi(, Sf~ál)so. Brazil (cull. A. lI'{lg,,~r s.".): G. Trichocf'mmll/
C"(/mJidr'lII. Gualem:lIu (D. Milldslllf'd, S./I.): 11. TridUJct'ntrum dilllllll1,III. Costa Rica (F. P"puN" 20): 1. Tr;cho<:tmlru'" fi'.~cum.
Brazil (Seidel s.".). 8:trs • l cm.
O
•
185
PUPULl N- A REV ISION OF TR /CIfOCENrRUM
diinae inlo a unique Rodriglle1.ia elade (main ly
composed by Iwig-epiphyles). wilh Ihe unique excepdon of Trichocerllrum. Ihal Ihe aulhors consider, phylogenelically, an artificial genus. I hope
a rev ision of Trit:llOCelllrulll may conlribule lo
beUer undersland ils relalionships wilh Ihe main
corpus of Ihe O ncidii nae and l..ophiaris complex.
In spite of a cerlain number of common fealures.
Trichocentrum species in facl prcsenl very differenl Irails. and some of Ihese characlers muy prove
usefu l to recogni ze the occure nee of several. differe nl natural groups within Ihe genus.
Characters used in groups and species
delimitation
Habit (l lId size-Spec ies of Trichocelltm m
have a generally simi lar habit and are extremely
variable with rcspect to plant size. Gross differences in size do exist among differenl groups of
laxa (Le.. planls of Tric/lOce fllmm CllllditJllm
group are generall y smallcr Ihan laxa perlaining
10 Tric/IQCelltrllm f llSC1II1I group). However. Ihe
size of individual plants can be excessively infl uenced by environmental elements. and the grcat
variation wilhin populations results in size bei ng
of no practical utilily for laxonomic purposes.
Roo/s-Trichocell/rum species share Ihe same
basic rOOI Slruclure which is Iypical of Ihe subIribe. with long pericyclal cells. a broad-celled
conex o a thin layer of endodermis cells and 3-4
layers of vclamen cell s. Major varimi ons are associaled 10 Ihe Ihickness of the conexo rcsulting
in different rool diameters, Ihat can vary from 0.8
10 1.9 mm. Such a difference. however. is chieny
consistent wi thin well defi ned spccies complexes,
and the charucler has bul a li u le praclicul value
in delimiting spcc ies.
Pselldoblllbs-Triehocelllrum pseudobubs are
very small. usually less Ihan 0.5 cm in diameter.
and vary in shapc from subg lobose 10 ell ipticalovale to cy lindrical. The lalter Iype is gencrally
longer. being sometimes 1.0 cm long. Varialions
in pseudobulb size, however, appcar 10 be consislenlly associaled to Ihe dimension of the apical
leaf. Ihus resuhing of no vulue in species delimitati on.
Leaves-Leaf shape, coloration and thickness
are variable characlers in most of Trichocelllrum
spccies. As a general ru le. T. tigrirwm and some
other species show leaf adaplalion 10 eliophylism
(Le., red-pu rple pygrnenlalion) or hydric Slress,
186
whereas leaves of mOSI of the species are Iypical
of shudow-growing, moist dependent epiphytes.
Although such differences have been used 10 delimil species (Reichenbach, 1876; Rolfe. 1893;
Beckner. 1961 ; Schweinfurth. 1945). variations in
these characters are mainly dependent on environmenlal elements and may also vary wilhin individuals.
Jlljloreseellee- Mosl Trie/weel//mlll spec ies
present successive inflorescences producing Ihree
10 many fl owers over a long periodo In many
cases, if pollination does not take place along Ihe
princi pal ax is of the innorescence, racemes may
freely branch bccoming paniculate. but T. calo·
cera.f also show primarily panicul ale inflorescence. Singlc-fl owering inflorescences are not uncommon in South American laxa. Simultaneous.
Iwo-nowered, somelimes mulli-branched inflorescences can be found in sorne species comp1exes:
a mu lti fl owered and sirnul laneous inflorescence
Iype has been also recorded (Schwei nfurth. 1946).
Though generally ignored, innorescence type and
nu rnber of flowe rs are useful churacters 10 disti nguish species.
Ovary-Most of Trie/¡ocelllrtllll species possess terete to subclavate ovaries. with the eltception of Ihe spec ies elosely related wilh Tridw·
centmm plllchmm. Ihal can be easily distinguished from Tric/lOcentmm [lIscllm group on Ihe
basis of their unique lriquetrous ovary.
Sepllls ami pewls-Differences in shape and
size of the sepals and petals have been used in
diffe rem ialing species (Lindley. 1850; Reichenbach, 1883: Schlechter. 1923). Flowers of TridlOcelll rtlm generall y bear si milar sepals and pelal s.
varying in shape from elli ptic 10 linear-ell iptic to
elli ptic-obovale. Tepals vary in length from 5 mm
to about 30 mm. Spreading or cJosed sepals and
pelals permit 10 de limi t sorne laxa. Size differences umong species can be observed and uti lizcd
in delimiting spccies complexes, but the strong
depcndence of flowe r size on environmental elements reduces Ihe practical value of this character
for spccies dcl imitation.
ulbellum-S hape und ornalllenlation of Ihe labellu m are sorne of the most useful characters to
distinguish groups and species within Trichoct!n ·
tmm. Labellum morphology has been extensively
used in Ihe elassificalion of Ihe genus (Rcichenbach. 1876; Krlinzli n. 1897; Lindley, 1850; Schlechter. 1923; Garay. 1970; Pupulin. 1994 ; Pupulin
PUPULlN- A REVISION OF TRICHOCENfRUM
& Mora-Retana, 1994), and differences among
taxa are sufficiently consistent to assume practical
value in species delimilation. General shape of the
lip, shape and number of the basal calli , presence
or absence of lateral lobes, degree of apical and
lateral margins of the labellum were used in {he
present analysis.
Spur- The length of the spur has been used to
di stingui sh si milar taxa ( Ro lfe, 1894 , 1913 ;
Teuscher, 1961 ). bUI {his $Ole characler proved lO
be rarely critic in distinguishing species pertainíng 10 the same group. Shape characters that
proved to be usefu! in deli mitation of species
complexes inelude !obulate, cylindric. and elavate
spurs. Also the indumentum of the intemal walls
of the spur may be utilized 10 distinguish groups
of taxa. On the contrary, number of spur lobes in
taxa presenting lobulate spurs has nOI practical
value in differentiating species. and many observalions confirmed the variability of this character
also within different flowers on the same inflorescence. Although some early workers emphasized the grade of spur sinuosity in long-spurred
laxa as a diagnostic charaCler (Lindley. 1843;
1854; Reichenbach, 1883), the great variability of
thi s character within popu lalions results of unpractical value for taxonomic work.
Column- Apan from a few cases (Reichenbach , 1854; 1869; 1881; 1883), morphological
characters of the column have been largely neglecled in delimiting species concepts within Trichocenlrum. Columnar characters contributed to
lhe delimilarion of some laxa in thi s treatment.
Trichocenlrum species generally share the same
basic structure of lhe gynoslem ium. with a short,
stou t, winged column and a broad sligma, bUl the
wings may show considerabl y differences between elosely related taxa. Useful characters include presence or absence of glandu lar appendages at the column apex , shape. color and margins
of the wings.
Amher-Indurnenturn lype of lhe operculum is
a consistenl character and it has been used in this
trealment to delimit species within sorne groups.
Pollillarium- A lthough generally ignored in
spccies descriptions. pollinarium characters may
prove to be useful in del imiting differenl taxa
within the genus. The main differences in pollinarium in volve stipe morphology and the degree
of movement after it is removed from the column.
These inelude the grade of reflexion of distal mar187
gins and the placement of pollinia in various posilions. Such characters were not used here to dislinguish different taxa because of the limited information available on the pollinarium morphology of many Trichocemrum species.
TAXONOMY
Trichocenlrum Poepp. & Endl .• Nov. Gen.
Spec. PI. 2: 11 , t. 115. 1836.
Acoidium Lindl., Edwards's BOl. Reg. 23:
sub l. 195 1. 1837.
Orchis Ruiz & Pavón, in Herb. Real Jard.
Bot. Madrid , Ms., anfe 1788.
Perennial herbs, epiphytic. with vegetati ve
buds developing from lhe base of the previous
growth on a reduced rhizome. Pseudobulbs more
or less inconspicuous, ovoid or rounded to cyli ndric. generally concealed by scarious braCIS.
RoolS filiform or fleshy, glabrous. somewhat
branched, arising from the base of the new pseudobulb. whitish with green apex. Leaves single,
only occasionally in pairs. articulate al the apex
of the pseudobulb, fleshy to coriaceous, usually
small, suberect to arcuate or pendent. persistent;
varying in outline from ovate to oblong-elliptic to
ligulate, usually widest in basal half. mor or less
gradually narrowing to a sessi le base. InHorescence basal or terminal, simple or racemose,
somewhat branched, producing I 10 many fl owers. often successively on a shon rhachís, arising
from the base or near the insenion of the leaf of
the last pseudobu lb; pedunele terete suberect,
arching or pendenl with two or more lanceolate
acute bracts; rhachis usually fractiflex. Flowers
often large, showy and sometimes fragran!. Sepals and petals free, subsimilar, usually spreading, wilh acute to emarginate apex . Llp adnale 10
lhe base of the column, suberect. simple or obscurely three-Iobed, concave, fiat or convex, with
or wilhoul calli or ridges near the base, producing
wilh the column base a slender or gibbous or clavale. shon or elongate spur. Column shon. stoul.
adnate lo the lip, auriculate or wínged aboye,
without a foo!. Clinandrium commonly shallow,
with emire margins. Anther cucullate to emispherical , wilh 2 cells, glabrate 10 hirsu{e. Pollinia
2. nOlched to complanale, on a triangular more or
less elongate slipe often with refl exed margins;
viscidium peltate or ovoid.
D1 STRIBUTIQN : AboUl 23 species wide-
PUPULl N- A REVISION
or TR/CIIOCENTRUM
a natural group, Besides Tr;clu)(:entrum cllpis/ra-
spread from southe ro Mex ico through Central
America southward to Brazi l, Boli via, and Peru.
111m. the range of whic h ex.(c nds southward 10 Co-
lombia and Venezuela. aH the remaining species
in th is gmup are restrie ted to Central America
north of the Panama Canal. The greater concenlralion of species can be found in Cosla Rica:
only one species rcaches Ihe northem slates of
Veracruz and Guerrero in Mexico. This panero
suggests central Costa Rica is Ihe cenler of di sIribulion and the possibility of sympalric specialion in the group (Fig. 3),
Key to Ihe Groups of Trichocelltrum
la. Spur gibbous. 2-4-lobed. or clavale ......... ... 2
2a. Spur short, Icss Ihan S mm long. 2-4 lobed; lip
elliptie-Q\'alc . . . . . . . . . . . . . T. nltldidum group
2b. Spur ionser lhan 10 mm, clavale: lip paOOunlle
........•..•.•..•.•..... T. hM8<,i group
. . . . . . . .. 3
lb. Spur slendcr, cOl1ic or lubular ..
33. Lip wilh IWO Imcral. creel. (a!cme l<:>bes n-ear lhe
base
. . . . . . . . . . . . . . . . . . . T. pfm'ií group
3b. Lip wilhoul Imeru!. ereel. fa!cale lobes near lhe
base
.. .....
4
4a. Spur Ion Ser or cqualling lhe lip, tlexuous 10
unci nale . . . . . . . . . . . . . . ... . ...
5
5a. Ovar), Inquelrous; dis<: Ihickl)' bicarinale
......... T. ¡m/chrum group
Sb. Ovar)' nOI lriquclrous; dis<: will!. IWO 10
four slcndcr earines
T. fiucum group
4b. Spur shOrler lhan lip. linear
6
6a. Li p eoncave. slronSI)' narrowed above;
nowers campa nulalc, produced on suceessive in tlorescence ........ .
T. brachyculls group
ób. Lip nal 10 slighll)' convelt, Wilh di lated
lamina; nowers sprcading on 1- 2-nowered. simullaneous innorescences
T. ligrinum group
Key lo 'he s pecies of Ihe
Trichocelltrllm calldidlllll gl"oup
13. Amher cap glubrous . . . . . . . . . . . . . . . T. c{¡/oceras
lb. Anl her cup papi llose 0 1 hirsu te
2
2a. Column wi ngs dccurrclll, obtuse. will!. inlrorse
apexes .......
. ..... T. CfJJlllricellJe
2b. Colu mn wings ascending or porreel, aCUle. wilh
3
some.... hal revolule apex es ...........
3u. Li p narrow in Ihe middle; nowers 5mall (sepals and pelals 10 5-6 mm long); innorescence eree! 10 palenl ...... , ... T. brelresií
3b. Lip elliptic ovale 0 1 rhombic; nowers medium 10 large (sepals uOO pelals tonger Ihan 1S
mm); innoresccnce peOOule .......
4
4a. Up shorter Ihun sepals, acule 10 obIuse
.............. T. CflpiS/rfl/um
4b. Up longer Ih:m sepals, reluse al apelt
S
Sao Up obovale: oolumn wings cmirc
. . . . . . , . , . , . , ..... T. ronJidum
Sb. Lip broadl ), rhombie: colu mn wings
erose UI lhe apel! ... T. cJlllbig/ossu",
Relationships belwccn TricllOcemmm groups
suggesled here are summarized in Fig. 2.
Tric/lOcelltrum ca"didum group
Thi s group inel udes six spccies presenting a
gibbous. saecatc. 2-4-lobcd spur, namely T. brenes;; Sc hhr.. T. caloce ras Endres & Rchb.f., T.
calldidllm Lindl.. T. l:a/Jisrrtllll m Linden &
Rchb.r.. T. cO.fraricell.fe Mora-Retana & Pupu lin ,
and T. c)'mbig/os.fllm Pupulin. Apart fmm the
rather obscurc T. brenes;;, known only from a
copy of Schlechter's druwi ng or the typc, all the
spccies wilhi n Ihe group present a delicate. semi hyal ine texture, with simi lar sepals and pctals
varyi ng in colour from wh ite to greeni sh-white or
browni sh ye llow. The lip is Hat or coneave, without any callosi ly or wilh IwO obscure thickened
kcels near the base, Furthcrmore. all the taxa invariably presenl filiform mots, a successive inHorescence and a gibbous, short, 2-4-lobcd spur.
The stipe is mobile in alllhe spec ies, and strongly
reHecls its lateral distal margins after a few seconds fmm re moval. Pollinia are e ntire. and Ihey
do not overl ap aftcr reflection of the slipe. AII the
species are morphologically very well differenliated from the main body of the genus. a nd their
nalural distribution confirms that they constitute
T. brenes;; Schltr.. Re/Jert. S/)(!c. Nov. Reg/li Veg,
Be;/¡. 19: 248. 1923. TYPE: COSTA RICA. AI-
ajuela: San Pedro de San Ram6n, Sep!. 192 1, A.
Brenes 116 (Hololypc. B, destmyed: Lectotype,
AMES).
DISTR IBUTlON: Endemic 10 Cosla Rica.
REPR ESENTATI VE S PEC IMENS: COSTA
RICA. Alajuela: San Je r6nimo, C. Werc:k1é J39
( 8. destroyed). Wilhoul locality. Nel'ermatlll S. /I.
(B. destroyed).
T. caloceras Endrcs & Rchb.f.. Card. C/¡rQn.
1257.187 1. in/exllI. TYPE: COSTA RICA. With·
out specific locali ly. 1867, Emires 5.1/. (Typc not
localed: neotypc. herc designated. COSTA RICA.
Puntarenas: Coto Bros, Las Cruces, F , PllplI/;1/ &:
D. Castelfral/co 1, USJ).
DlSTR IBUTlON : Costa Rica and Pana ma.
REPR ESENTAT IVE SPEC IMENS: COSTA
RICA . San José: Perez ZcledÓn. San Juan de San
Isidro. F. PII/mli/l JO (USJ); Alto de San Juan, F.
Puplllill &: D,E, M ora-Re/lIIllI 186 and 187 (USJ).
188
PUPULl N-A REVISION OF TRICHOCENTRUM
Group
Characteristic
flowershape
labellum
Spur
shape
.hape
Catldidum
~
,.......
Hoegei
Plavil
Fig. 2.
....... ,
.... ...
~
D
~
Q(~~~
(}
The subdivision of TrichQunlrum.
189
(~
t:'?h
\@1
"
.- . .
@
"
"
(.j{.)
~
PUPULlN- A REV ISION OF TRfCIfOCt:NTIWM
•
','"
"'.
/'
• 'Os ,
*• T.T. c:aIocItras
bnlne$ii
eT.canddum
l>. T. capistratum
oTo eostarieltnM
... T. eymbiglossum
Fig. 3.
Distribution of Ihe speeies of Trichoct!nlru'" cOtulidum group.
Campana, V. Hellbuiek 528 (SEL). GUATEMALA. Quetzaltenango, Ch uikabal , Lewis 22/;
Guanagasapa, Finca Santa María Buena Visla, o.
Mitre/Maedl S.n. (Herb. Pupulin). MEX ICO. Veraeruz: beyo nd Bastioná l, R. Dre.\·s/er 4446
(FLAS); Tezonapa, Hacie nda La Unión, Nage/ &
Juan 4/75 (AMES). Chiapas: Soconusco, above
Huixtla, Nagel 4836 (AMES).
Puntarenas: Coto Brus, Las Cruces, F. Pupulin &
D. Ca.welfraneo J (USJ). PANAMA. Chiriquí: between Volcán and Concepción, C. Luer, A. Luer,
R. Dress/er & NH. William.l" /352 (SEL); C.
Luer, J. Luer & H. Butcher 9280 (SEL); H.P.
Buteher 652 and 653 (SEL).
T. candidum Lindl. , Edwards's Bot. Reg. 29:
mi se. 9. 1843. TYPE: GUATEMALA. Wilhout
specific 10calilY, Skinner S.f!. (Ho lolype, K).
Fig.4.
T. albiftorum Rolfe, Bull. Mise. Inform. 336.
1893. TYPE: MEXICO. Veracruz: withoul 10cality, Fink .~. n. (Hololype, K).
DISTRIBUTION: Mexico, Guatemala, El Salvador.
REPRESENTATlVE SPECIM ENS: EL SALVADOR. Naranjal Deininger, near San Salvador,
F. Hamer s.n. (Herb. Hamer); Hacienda Las Lajas, Volcán de Izealco, F. Hall/er 72 (AMES,
SEL); Cerro Monteeri sto, F. Hamer UI.; Cerro
T. cap;slratum Linden & Rchb.f., Gard. C/mm.
1257. 187 1. TYPE: COSTA RICA. Withoul spedile locali lY, Wa lli.~ .~.n. (Hololype, W).
T. panamense Rolre, Bull. Müc. /nform. 341.
19 13. TYPE: PANAMA . On bush-covered hills
easl of the Panama Canal, 255 m, 9°10' N,
79 0 41' W, \909, LJ. Lip.w:omb s.n. (Holotype,
K).
T. pusillum Lehm. in Herb. Kew., M.s.
DISTRIBUTION: Costa Rica, Panama, Colombia, and Venezuela.
REPRESENTATlVE SPECIMENS: COSTA
190
PUPULlN-A REVISION OF TRJCHOCENTRUM
5mm
5cm
2 cm
1 cm
""
1I
", ",::,'
J
3cm
3mm
Fig. 4. Trichocemrum candidum Lindl. VOllchcr: GUATEMALA. Guanagao;apa. Finca SaRta Mar(a Buena Vista.
s.n. (Hcrb. Pupu l in).
191
o.
Miuelsflledr
PUPULl N- A REV ISION OF TRICHOCENTRUM
RICA. San José: Pérez ZeledÓn. Alfombra de San
Juan. F. Pupulin 3 (USJ); Las Nubes de Quizarrá.
F. Pupulin & 1. Cambrollero 4 (USJ); Rivas. F.
Pupuli1l & 1. Cambronero 7 (USJ). Pumarcnas:
Cañas de Buenos Aires. J. Cambronero S.1I.
(USJ). PANAMA. Upper Chagres Ri ver. Powe//
419 (AMES); Lake Gatún . Barro Colo rado Is ..
J.o. Ackerman 13 78 (S EL). Coc lé: near Ihe edge
of El Valle de Antón. R. Dre.~.fler. e Luer. l. Luer
& P. Taylor 766 (SEL); lower pan of El Valle de
Antón. Hulller & A lfen 384 (AMES); near Achiote. R. Dress/er 3 / 46 (FLAS); belween Cerro
Azul and Cerro Jefe. R. Oreu/er 3263 (FLAS);
e. U/er. l. Luer & H. BUlcher 9/ 70 (SEL); El
Valle de Antón, 1. ClImbrollero .f.n. (Hcrb. Pupulin); lA F01l.feca .1'. 11 . (He rb. Pupulin). Withou t
specific localily. R. Oreu/er 2832 (SEL). COLOMBIA. Boyaca: El Humbo, w wrance 557
(AMES); Anlioqu ia: Fro ntino. near Dabeiba. Lehmallll 4633 ( K, AMES); Le. Vieira .f.II. (He rb.
Pupulin); San Carlos, G. E,H:obar 487 (A MES).
VEN EZUELA. Zulia: Cajime ra, Perijá. e. Garcia
E. .1'. 11. ( K ); Coló n, Río Catatumbo, D. Palrzek
S. /I.; Sierra de Perijá. Río Yasa. Sreyermllrk &
Femllm/ez 99625 (AMES).
T. CQstaricellse Mora- Rctana & Pupulin. Se/byarw 15: 94. 1994. TYPE: COSTA RICA. Alajue-
•••
. T. ~i
Fig. j .
la: San Carlos pl ain , near Ciudad Quesada, 1987.
Ho rich .UI. (Ho lo typc. USJ).
DISTRIB UTION : Ende mic to Costa Rica.
REPRESENTAT IVE S PEC IM ENS : COSTA
RICA. Alajuela: San Juan de San Ramón. 3 1 Oc!.
1989, F. PuplIlin & M, Flore.f /7 (USJ). Prov, of
Can ago: Turrialba : La Se lva, F. PI/pu fin & O,E.
Mo ra -Retalla 229 and 230 (USJ).
Oistribl.nion of TrichOCfnlrom
h~g~i.
Trie/wcelllrum hoegei group
e H.
This group ineludes a single species endemic
Mexico, ¡he spur of which is not conic or tubular, but elavate: Tricll()cel/trwn hoegei Rchb.f.
It represents an isolated tuon, seemingly witho ut
any close relati ve. The spec ies presents a subquadratc lip, with two small basal calli. surro unded by purple red necta r·gu ides, Its pollinarium
shows ¡he shorter sli pe of the enlire genus. with
re Hected distal margins. Pyriform, complanate
pollinia are wide, enlire, and do not overlap after
reHection of (he sri pe, Trichocentrum hoege; is
the o nly known species to have a elavate spur,
and its iso lated di stribution in Mexico (Fig. 5)
seems to support the hypothesis of a n early segregalion of this taxon from the main complex of
the genus.
10
T. cymbiglossum Pupulin , Lindleyana 9: 5 1.
1994. TY PE: COSTA RICA. Alajuela: San Carlos. epi phytic o n tru nks along Laguna Bosque
Alegre. F. P//lmlil/ 5 (Ho lotype. USJ ; lsotype.
Herb. Pupulin).
D1STRIB UTION: Endcmic 10 Costa Ri ca.
REPRESENTAT IVE S PECIM ENS: COSTA
RICA. Alajuela: San Carlos, Laguna Bosq ue Alegre. F. PI/pI/fin 5 bü (He rb. Pupulin); Reserva de
Juan Castro Blanco, A. Herrera S. /I. (USJ). Cartago: Turria lba, Tres Equis, 30 Oc!. 1989, F. PI/- • T. hoegei Rchb.f. , Gard. Chron. n.s. 16: 717.
pu/in & M. Flore.f 12 (USJ): Río Estrell a, near
188 1. TYPE: MEXICO. Veracruz: near Cordoba,
San Isidro de Tejar, 14 Nov. 1989. F. PU/Jlllill &
Hoege .1'. /1. (Ho lotype, W; llIustration in Xellia
A. Flores 18 (USJ): Palo mos de Santa Teresita,
O reh. 3: 69, 1. 234. Fi g. 6. 1890).
F. Pllpulill 195 (Herb. Pupu lin),
D1STRIBUTION : Endemic to Mexico.
192
PUPULl N-A REVISION OF TRICHOCENTRUM
5mm
1 cm
Fig.6.
Trichocemrum hoege; Rchb.f. Vouchcr: MEXICO. Ouaca: Rlo Salado. M.A. SolO 4324 (Hcrb. Pupulin).
193
PUPULlN- A REVISION OF TRICHOCE.NTRUM
..._~'~
~/'
/
. . ...~ j
' ..,--
\
.••..
~
.a. T. cfanlhum
.T. aStr.UenM
eT. plavii
Fig.7.
Distribulion of the species of Trichocenrrum pfav;; gTOl.lp.
trum dianthum exhale a slrong, herbaeeous perfume , that may be associaled with sorne special
pollination syndrome. lt is also likely Iha! the
struClure of Ihe papillose spur plays a fundamen tal role in pollinalion of Ihese taxa. The pollinators of (hese three species, however. are slill un known.
REPRESENTATlVE SPECIMENS: MEXICO.
Guerrero: Teotepec system, MI. Piloncillo, Nagel
/902 (AMES). Oaxaca: Río Salado, in oak rorest,
1400 m, M.A. Soto 4324 (Herb. Pupulin). Vemcruz : Bastionál, R.L. Dressler .~ub Hagsater
4452a (Herb. Pupulin).
Trichocenlrum pfavii group
Key to the species of Trichocentrum pfavii
Thi s group also appears 10 have originated in
group
Cenlml America, and ils species are limited to
central and soulhem Cosla Rica and the Chiriquí
la. Spur more Ihan 5 mm lon g; column wings with longitudinal brown stri pes
T. dianth ..m
province in nor1hem Panama (Fig. 7). Trichocenlb. Spur less Ihan 5 mm long; col umn wings whi lc wilh
trum pfavii Rchb.f. , T dianthum Pupulin & Morabrown SpolS al margi ns ... . . . . . ........ . .
2
Relana, and T e.slrellenu Pupulin & J.B. Garda
2a. Lip nabcllale. wi lh very crisped apex; latcral
lobes of the lip di gilale ........ . . . . . T. pfa"ii
are Ihe only three species wilhin the genus pre2b. Lip obovate. f1eshy, wilh plane apex: latcrall obcs
senling short, ereel lobes near Ihe base of the lip,
o r Ihe lip shon and roundcd .......T. U lre/lense
such as papillose-hirsute internal walls of the
spur. Due lo their vegelalive and floral morphol - • T. dio,nthum Pupulin & Mora-Retana, Selbyana
15: 90. 1994. TYPE: COSTA RICA . San José:
ogy these species are more similar to each olher
Perez Zeledón, Las Nubes de Qui zarrá. 1000 m,
than to any olher taxon in the genus. Trichocen194
PUPULlN- A REVISION OF TRICHOCENTRUM
epiphylic on short lrecs along a Hule river. J.
Cambronero .~. n. (Hololype. USJ).
DISTRIBUTION: Endemic lO Cosla Rica.
REPRESENTATIVE SPEC IM ENS: COSTA
RICA. San José: Allo de San Juan. F. PupuUn &
W. Forue!:a 20 (Herb. Pupulin); F. Pupulin & w.
FOl/seca 22 (USJ. Herb. Pupulin); Slopes of Cerro
de Las Vueltas. A. Herrera S. /I. (USJ). Wilhoul
localily. M . Flores s.n. (USJ).
T. estrellense Pupulin & J.B. Garda sp. nov.
TYPE: COSTA RICA. Cartago: El Guarco, in
vallcys under Palo Verde, epiphYlic in shadow on
lall trees along a minor tributary of Rfo Reventazón, 1250 m. 30 Apr. 1992, F. Pupu/in 209 (Holotype. USJ; lsolype. Herb. Pupulin). Fig. 8.
Species Trichocentro pfavii similis, sed labello
minore marginibus planis, lobulis lateralibus abbreviatis rotundali s, alis cotumnae abbreviatis
marginibus laceralis et inflorescenlia heterocronica differt.
Planl epiphytic, cespitose. pendent. wilh abbrevialcd rhi zome. RoolS filiform . flexuous, glabrous, silvcry white with grecn apcx. Pseudobulb
cylindric, up to 10 mm long, 5 mm wide, monophyllous. Lear lincar-clliplic 10 elliplic-oblong.
obtuse 10 rctuse. narrowcd 10 a conduplicate. sessile base. lighl grecn. 8- 12 cm long (10 20 cm),
2.5- 3 cm wide. InHorescence laleral. basal, pendenl. successive. sometime branched al the base,
3- to many-fl owered: pedunde lerete. 4 to 11.5
cm long. concealed by 2- 3 concave. ovale bracts.
Ovüry linear-davale, aboul 3 cm long including
Ihe pcdicel. Flowers rather large for the genus,
with free sepals and pelals. ivory white with a
narrow chcstnut brown blotch at the first Ihird of
their length and white lip. marked near the base
by two rose-purple bars. Dorsal sepal obovateelli ptic. obtuse to reluse, slightly coneave toward
the apcx, to 16 mm long. 8.5 mm wide. Lateral
sepals obliquely elfiptic. obtuse to retuse, with a
short. rounded apicule. subcarinale towards the
apcx and slightly concave, 10 16 mm long. 6.5
mm wide. Petals linear-el1 iplic . Obluse, sli ghtly
concave lowards Ihe apex, 16 mm long, 6.5 mm
wide. Lip obovate-subpandurate. adnale 10 the
column. 2 1 mm long. wilh a short. linear claw 2
mm long, with two broad, subquadrate. pubescenl
lateral lobes at the base, expanding abruptly to
the subpandurate blade. 16 mm long. 15.5 mm
195
wide, dceply emargi nate in front . with plane, entire margins. producing wi th Ihe co[umn base a
very short, blunt. saccate spur. 2.6 mm long. 3.5
mm wide. Column short, stOUI. wilhout a foot, 3
mm long, with a pair of short, erect, subdolabriform, lacerale wings, spotted and blolched with
brown along the margins. Anther white. cucullate, papillose. Pollinia 2. pyriform, rugu lose. on
a short obtriangu lar slipe; viscidium elliplic.
brown when fresh.
ETYMOLOOY: Named from La Estrella, lhe
region on northwestem slopes of the Cordillera
de Talamanca. where the species was first found.
DISTR1BUTION: Endemic 10 Costa Rica.
REPRESENTATlVE SPECIMENS: COSTA
RICA. Cartago: La Estrella. CH. Umke.wer s.n"
9 Jan. 1923 (AMES); El Guareo, bajo Palo Verde.
F. Pupulin 205 and 208 (Herb. Pupulin); Agua
Caliente, J.M. Quiroz S.II. (ftde J.B. Garda). Limón: Siqufrres. Bajos del Tigre. 450 m. L.. Acosta
& F. de Acosta .f.n. (ftde J.B. Garera).
Tricllocentrum estrel/ense inhabits submontane,
evergreen, tropical rain fore~;(s of the Atlantic
slopes of the Cordi llera de Talamanca and Cordi llera Central in Costa Rica. where the planls of this
spccies grow on shady branches and lrunks covered by live moss, mainly over slrcams. The cJ imale of the high basin of Río Reventazón is wel,
temperate. wi th a moderale water deficit (Herrera,
1985). Minimum and maximum temperalure ay·
erage at La Estrella region are 18 and 2 1 C. respectively. Flowering generally begins in carly
summer, and many flowers are produced successively from June to Septembcr. Tricllocentrum esIrellefue was first collectcd in January 1923 by
C.H. Lankesler near La Estrella. in Ihe Province of
Cartago. and a specimen was senl lO Oakes Ames
who idemificd it as T. pfavii (AMES 26 162). Another anonimous collection from lhe same area was
ilIustrated in a delicate and delightful walereolor
by Rafael Lucas Rodríguez Caballero. This plate.
togelher with many other botanica/ dmwings o(
Costa Rican orehids by Rodríguez. was posthumously published in 1986 ac; T. pfavii (Rodríguez
Caballero el al., 1986). Due to the variable regional
form s of T. pfavii (Pupu/in & Mora· Retana. 1994)
il is nOI surpri sing thal herbarium malerial of T.
e.wrellense was overlooked for over 70 years. [n
fact. most of Ihe morphological differences between the two laxa are better observed in living
material. and !he characteristic successive inflores-
PUPULlN- A REVIS ION OF TRIC HOCENTRUM
5mm
3mm
3 cm
3cm
Fig. 8. Trichocemmm eslrelleltSl' Pupulio & J.B. Carera. Vouchcr: COSTA RICA. Canago: El Guareo. io valleys undcr Palo
Verde, F. Pup"lin 2Q9 (USJ ).
196
PUPULlN-A REVISION OF TRICHOCENTRUM
cence, unique among the species close 10 T. pfavii,
may be nOliced only when pla nls are kepl in cu ltivalion. Joaquin B. Gareía fi rst poi med OUI the
difference between planls native of La Estrella area
and lhe typical fonn of T. pfavii. In April 1992 I
had Ihe opponunity to confinn Ihe spccific status
of T. eSlrellelue after a visit to Ihe Valle del Guarco. in Ihe high basin of Río Revemazón, wherc
large colonies of Ihis species were found. Wilhin
Ihe genus, T. estrelltmse is so far the species with
longer leaves: a leaf of lhe type specimen was 20
cm long.
T. pfavii Rchb.f., Gard. Chroll. n.s. 16: 70. 1881.
TYPE: PANAM A. Chiriquí: without spccific 10cality, 19 Nov. 1880, Pfau 60 (HololYpe, W).
Fi g. 9.
T. ¡Jfavü varo ZOl/ale Rchb.f.. Gard. C/¡ roll.
n.s. 19: 44. 1883. TYPE: COSTA RICA. San
José: Boca de DOla, along Ihe c racks on the
Irunks of young Irces, Endre.~ .~'. I/ . (Holotype,
RL Dre.uler 3760 (FLAS); near Volcán, A. Dopp
S. II .
(SEL).
Trichocentrum pulchrum group
The dislribulion of the Ihree species in this
group is li mited 10 Ihe Andes from Venezuela and
Colombia lO Peru (Fi g. 10). These spec ies can be
casily recognized by Iheir Iriquelrous ovaries, a
unique characler wi lhin Ihe genus. Furthermore,
Ihey presenl two very Ihickened keels ari sing
from the base of the disco diverging al apex and
somcwhat inlensc ly pubcrule nt. The overall mQrphological similarily among Ihe Ihree species suggests a common ancestor for Ihe group. T he flow e rs nre produced on si ngle-f1owered, archillg lo
pcndcllI inflorescences. Tric/lOcemrum pulcllrum
is one of Ihe mosl widespread species of Ihe geIlU S: il is known from Venezuela lo Peru.
Key lo Ihe species of Trichocentrum pulchrum
group
h. Spu r Icss lhan twice as long as the lip ... T.IJIIlchrum
........ 2
la. Li p nabell ate. with a broad. dark purple blotch on
lhe d isc . . . . . . . . . . . . . . . . . . . . . T. brandlin ..
2b. Lip OOovale. wilh a yc llow blotch 011 lhe disc ..
T. fOllgicufcuro/U1II
W ).
J b. Spu r more than Iwice as long as the lip
T. ,mullder.fiWIIII11 Endres & Rc hb.f. in Reic henbach He rb., M.f.
T. saundersü Endrcs & Rchb.f. in Reic henbach Herb., Ms.
T. ZOl/a/e Rchb.f. in Reichenbach He rb.. Ms.
T. pfav;; f. al/mm He nderson in SEL, Ms.
D1STRIB UTION: Costa Rica and Panama.
REPRESENTATlVE SPECIMENS: COSTA
RICA. San José: Boca de Dota, Elldres S. II . (W);
San Marcos, road lO San Joaquin , F. Pllplllill &
M . Flo res 6 (USJ ): same localilY, F. PI/pufin 45
(USJ. Herb. Pupulin); Pérez Ze ledÓn. Rivas, F.
Pllpllfil/ & l . Ca mbronero 8 (USJ); Alto de San
Juan, F. Pupu/;n & W. Fonseca 2/ (USJ): vici nity
of El General. Skurch 220 1 (AMES). Prov. of
Cartago: Pejival1e, E"dres s. n. (W); El Muñeco,
south of Navarro. Stal/d/e)' 33735 (A MES): S/andley & To rre.f 5 1/ /4 (A MES); Slo rk 2734
(AMES); Orosi, Va/erio 2696 (AMES). Puntare nas: Coto Brus, Agua Bue na, F. PII/Jlllill 1/
(USJ); lA. FOII.feca .f.lI. (USJ): Campo Dos, F.
Pupulill & 1. Cambrollero 16 (USJ); Las Cruces.
F. Pupu/in & D. Casrelfrom:o 2 (USJ): Jardín Botánico Wilson, M . Cra)'um 9268 (MO). Without
locality: Elldre.f .~.I/. (W); C H. Lal/ke.wer .~. II . (K).
PANAMA. Chiriquí: bclow Volcán, "Cordille ra".
C Lue r, l . Luer, H. Burcher & A. Maduro 9319A
(SEL); vicinity of Volcán. No N. Wifffiams sub
T. bralld/Me Kraenzl., Bull. Herb. Bo is.~. 5: 109.
1897. TYPE: COLOMB IA. WithoullocalilY. Bar·
bey .~. 1l. (HololYpe nOI located).
T. verrucifemlll Schllr., Repert. S,Jec. Nov.
Reglli Veg. 7: 184. 1920. TYPE: COLOMBIA .
Without locality (Holotype B. deslroyed. drawing, Re¡Je rr. S¡Jec. Nov. Regl/i Veg. Deih. 57: t.
253. 1929).
DISTRIBUTION : Ende mic 10 Colombia.
REPRESENTATIVE SPECIMENS. COLOMBIA. Dept. of Risaralda, near Apia, CL Vieira
S. II . (Herb. Pupul in): Pueblo Rico, G. Escobar 70,
9 / and 554 (AMES). Depl. ofCauca, cult. A. Lehm(lll11 .mb C Luer 2969 (SEL).
T. IOllgicakara/um Rolfe, Orel!. Rev. 4: 260.
1896. TYPE: COLOMBIA . wilhout speci fi c 10cali ly. F.C /..ehmwlI! .f. lI. (Hololype, K).
DISTRIB UTION: Colombia and Ecuador.
REPRESENTATIVE SPECIMENS: COLOMBIA. Wilhout locality. Lehmalll1 3330 ( K). /..elllIIallll ex ID. Briell .f. II . (K). ECUADOR . Azuay:
Cuenca, Lehmolll1 8379 (AMES). Napa: near
Tena, C H. Dodsol/ SoI/. ( 1987). Zamora-Chi nchipe. ncar Zamora. C H. Dodsoll /7828 (RPSC).
197
PUPULIN- A REV ISION OF TlUCHOCt,WTRUM
5mm
,
3cm
5mm
Fig, 9.
5mm
Trichocemrum pfavii Rchb.f. Voocher: COSTA RI CA. Pumarenas: Las Cruces, f : PU(JUlin &. D. Cuslelfrum;a 2 (US}),
198
PUPULlN- A REVIS ION OF TRIC HOCI;"lffRUM
•
•
"
••
<"
~:-_ ......-"(
r
,-
l"
.
"
!
"
Sehlim 4/ (as T. speciosum, K. P); Salldeman S. II .
(K); Aspasica, ex Hort. Linden (SR). Rfohacha:
Sierra Nevada de Santa Marta, Lillden 1666 (type
of T. /IIaellla/llm, K. SR, P. W). VENEZUELA .
Lara: Yacambrf, J. V. Stope/o .~.II. Zuli a: Perijá,
Río Omira- kuná, Steyermark & DIII/slerville
/056/0 (AMES). Mérida: near La Carbonera. E.
Fo/dats VI. Trujillo: near Boconó. G.c.K. & E.
Ollmterville 551. ECUADOR. Garuma les. Río
Paute, P. Alldreetta 3302 (SEL). Pastaza: Mera,
E. Asp/lllld 194/5 (AMES. K). South of Laja,
O'A1essalldro 619 (MO). Zamora-Chi nc hi pe ;
Zumba, F. Pupll/il! 254, 255. 256, 260 and 263
(Herb. Pupu li n); O'A le.uandro 751 (RPSC); Cerón 138 /2 (QCNE); OOd.WlI /5798 (MO). PERU .
Junín: Tarma. D.E Bennell & C. H. Dot/so" 6773 (SEL); Cha nchamayo, aboye San Ramón, D.
Bellllett & A. Be,,"ett 0677-/-7; o. Bellnett &
C H. DotIsolI 677 (MO. AMES. UC).
Triehocentrum fu seum group
•
•
.. T. brarldliM
• T. Iongicalcan'lUJm
• T. pulehrum
Fig. 10. Dislribution of the spcdes of TrichoUlllrwn pulehr,,'" group.
T. pulcltrum Poepp. & Endl.. Nov. Ge/l. Sp. PI.
2: 1\. 1. 11 5. 1836. TYPE: PERU. Huánuco: near
Pampayacu, Poeppig S. /I. (Holotype. W). Fig. 11 .
Orehü punetala Ruiz & Pavón, in Herb.
Real Jard . Bol. Madrid, Ms.
T. /IIl/eu/atlllll Lind1. , Orel/id. Li/lden. 24.
1846. TYPE: COLOMB IA. Río Hacha. Sierra
de Santa Marta. alo ng the Río San Anlo nio
( 1300 m). 1844. Linden 1666 (HoloIYpe. K; Isolypes. BR. P. W).
T. .VJecioslIlI/ Sch lim. in He rb. Kew.. Ms.
DISTR IBUTION: Colombia. Venezuela. Ecuador. and Peru .
REPRESENTATIVE SPECIMENS : COLOMBIA. Cundinamarca: Bogotá. G. Ka/breyer S.II.
( K); o.~/Jilla - Hernandez 2/ / (AMES); La Mesa,
e/l/eS 98 (P) a nd 2859 (BR); Soulh of Bogolá,
near Guayabetal. R. E~eobar S. /I . (Herb. Pupulin);
Guayabctal. ex Hort. Colomborquídeas (Herb.
Pupulin); Villavicenc io. C Slllldellll/I/ S. II . (K); region Mera. Clae.~ S. II. (AMES). Ocana: La Mata.
The six species of this group are found from
Venezuela, through the Guianas. 10 southern Brazil , and from Ecuador to Bolivia and Peru along
the Andean Cordille ras (Fig. 12). Apart from T.
palldllrOtllm, the ft owers of these six species are
produced successively o n generally pendu lous inftorescences. The group shares the long. ft exuous.
slender spur and the re lalively complex calli al
the base of the \ip. two features I believe are derived wi thin Trichoeetllrum. The lip is generally
wide, subpandurate 10 pandurate in OUl line, sometimes wi lh brightly colored nectar guides toward
the e ntrance of the spur. Tho ugh no dala on the
polli nation of any of Ihe species in thi s group are
available, it is highly probable that the slender
elongale spur is someway involved in attracling
poll inators. Triehocentrum fll.sclI/ll Li ndl. is Ihe
most widely diSlribuled spec ies of Ihe group: it is
recorded from Venezuela through Ihe Guianas lo
Brazil and the Amazonian regions of Ecuador,
Boli via and Peru .
Key to the species of Triehocenlrmn fuseu",
group
la. Inflon:sccnce simuhancous ly 2-
5_nowercd
T. (Nm¡/urtllmll
t b. Innon:sccllCc suceessivc
. .. .. .. ..
2
211. Column wings lallCcolalc !O slrielly triangular .. 3
3a. Disc of lhe lip wi!h four kec ls ..
T. o/oo-coccinrum
199
!o
PUPULl N-Á REVI$IQN QF TRICHOCENTRUM
5mm
/);;
~
2mm
5mm
1cm
Fig. 11. TridlO,:eutr"m Imldm"" Pocpp. & Endl. Voucocr: COLOM BIA. Cundinamarca: Guayabelal. ex Hr>r(. Colomborquide(.l"
(Hcrb. PupuJin).
200
PUPULlN- A REVISION OF
o.
o
"
•
...
1..,0\
o
.. -
__ o
.-<..
'.
!
.. T.
albo~
• T. 1\.1_
O T. pMclmlturn
*.T.T. purpureum
rlCUMlm
6. T. lenuillorum
Fig. 12.
Dislri bulion of Ihe:
~pcdc:.~
of TrichQce",,,,,mju5c,,'"
groop.
3b. Disc o f !he lip wilh IWO keels .. T. " ,r,,!florum
2b. CoIumn wings broad. obtuse 10 truncatc al apex . . 4
4a. Disc of lhe: lip wilh IWO kee ls . . . T. rl'{:un'um
4b. Disc of !he lip wilh foor kee ls . . . . . . . .
5
5a. Spur aboul Iwice long Ihan lhe: Jip . . . .
T. ..'tIK""'¡
5b. Spur as long as Ihe: Iip o r shone:r ... . . 6
OO. Lip obovlllc·oblong ... T. pU'fmreum
6b. Lip subpandura!e: . .. .. , .. T. fiucum
T. albo-coccineum Linden . Belg. Ho rt. 15: 103.
1865. TYPE: BRAZIL. Amazonas: epiphytic in
forest along (he Rio Negro. Wallis S. /I. (Hololype.
W ).
T. albopllrpllreum Linden & Rc hb. f.. Gard.
Chron. 2 19. 1866. TYPE: based on (he same
type as T. albo·coccineum Linden (Holotype.
W).
T. albopurpureum varo striatum Linden &
Roo .. Lindenia 2: 77. 1. 85. 1866.
T. alboviolaceum Schltr., Die Orchjdetm 446.
1927, spha/m.
T. a maZOllicum Barb. ROOr.. Gell. Sp. Oreh.
Nov. 2: 140. 1882. TY PE: BRAZ IL. Villa Bella
20 1
TRICIfOCENTRUM
da Imperatriz, along Mac urany lake, Ba rhosa
Rodrigues S.II. (Holotype nOI located).
T. a/ropurlmreum Linden ex Rege1, Index
Se",. Hon. Petropo/. 79. 1868.
T. ionophtalmwfI Rchb.f.. Gard. Chroll. n.s.
6: lOO, 1876. TYPE: BRAZIL. cu/I. Ha",hurg
Bot. Gard. .~. ". (Holotype, W).
T. /eeanllm Rchb.f. . Flora 69:550. 1886.
TYPE: PERU (" Ex cord. occid. Am . aequat.") .
Ex hort. Lee (Hololype, W).
T. orthop/et:trorl Rchb.f. . Gord. Cli ron. n.s.
19: 562. 1883. TYPE: Origin unknown, ex
horl. Linde" .~. n. (Holotype. W).
T. IJOrpliyrjo Rchb.f.. 111. Hort. 3 1: 9, t. 508 .
1884. TYPE: Origin unknown, ex hon. Lil/{Jen
.~. II. (Holotypc. W).
DlSTR IB UTION: Brazil , Bolivia, and Peru .
REPRESENTATIVE SPECIMENS: BRAZ IL.
Wi thoul collection dala (BR). BOLIVIA. Santa
Cruz: Chiquitos. Robaré, Vasquez 95 1 (Herb.
Va sqllez janl/m ): Las Manos (AMES). PERU .
Amazonas: Bagu.. : Río Marañon. Hulc:hillSUII
1536 (A MES. K); La Peca. between Marañon and
Ulcubamba rivers. WOYlkowski 37027 (F); near
Bagua Chica. F/owards 48 (AMES). Huancabambao Lowii .1'.11 . (W). San Martrn : San Martín. Shapaja. D. Bennell & A. Belllum 15/8.
T. ¡useum Lindl .. Edward.f's Bot. Reg. 23: sub t.
1951. 1837. TYPE: incorrcctl y said 10 be from
Mexi co, c ulli valcd in the co ll ec tio n of Mr.
Knighl , Warmi"8 s.l1. (Holotype not localed;
drawing, K). Fig. 13.
AelJidiwlI /I/.w.:um Lindl. . Edll'ards's Bo/.
Reg. 23: su b 1. 1951. 1837. "0111. nI/d.
T. comllcopiae Linden & Rchb. f.. Gord.
C!mm. 266. 1866. TYPE: BRAZIL. Amazonas:
Río Negro. WaJlis S. II. (HololYpe, W ).
T. IUl r/ii Rolfe, BI/II. Mj.~c. ¡,,/orm. 395 .
1894. TYPE: VENEZUELA . Monagas: Maturín , c ult. Kew Bol. Gard . ex Han .~. ". (Holotype. K).
T. mattogro.\'.\·eflse Hochne. COII/III. Lill. Telegr., Bot. AmI. 5, part 1: 55. 1910. TYPE: BRAZIL. San Luiz de Cnceres. aloog (he Rio Paraguay. Hoehne .\'. 11. (S P, not seen).
T. pjl/ehi Lindl.. Gard. e hro" . 772. 1854.
TYPE: BRAZIL. Wilhoul specific locality, Pille/ .1'.11. (HoIOlYPC. K).
DlSTRIBUTION: Venezuela. Surinam. Brazi l,
Ecuador, Pero. and Boli via.
PUPULlN-A REVI$ION OF TRICHOCENTRUM
Smm
~-~
5 cm
3cm
1 cm
Fig. 1l .
Trichoc~nlrum ¡IISCllm
Smm
Lind l. Voocner: BRAZIL Bahia: IIIlCUS, CIIII. PIlPlllin u
202
~id~f
S.n. (Herb. Pupulin ).
PUPULlN-A REVIS ION OF TRICHOCENTR UM
REPR ESENTAT IVE SPEC IM ENS: VENE4
ZUELA. Bolívar: belween Tumercmo and El 00 4
rado, C.C. K. & E. Dunsrerville 409; north of
Tumeremo, C.c. K. & E. DllfI...rerville ....,1.: near
Ihe Angel Falls, C.C.K. & E. DlIlISfervilfe ."./1.
SURINAM . Brownsberg Mazaroni top, Determmm 154 and /63 (SEL). FRENCH GUIANA :
Monl Atachi Bacca, région de rInini. 7 km a r est
de Gobaya Sou la, 480 m, 53~ 55 ', 3 1 ~ 33', Cra1l
vilfe el al. (AY). BRAZIL: Amazonas: Río Negro.
without collcetor (BR); Wallis ."./1. (type or T.
cOmuCOIJ;ae. W). Bahía: Ilheus, cuJ¡. PU1J11U" ex
Seidel 5. 11. (Herb. Pupulin). Goiás: Inhumas, mil.
Pupll/il/ ex Seidel ."./1. (Herb. Pupulin). Pernan4
buco, wi lhout specific localilY, PjLwer S. II. (Herb.
Pupulin). Rio de Janei ro: along the Rio Paquequer, Neves AmlOnd 40 (B R); Rio de Janeiro,
Gaudic/umd 377 (P). Matto Grosso: San Lui z de
Caceres. l/oe/m e ."./1. (SP). Espirito Santo: Linhares4Mun -Callatine. Mlllford & Qaille Fmter
.".1/. (AMES). Without locality, l/on. Sal/der (K.
BR). ECUADOR: Zamora-Chinchipe: Paqui sha.
F. Pllfmlil/ 272 and 273 (Herb. Pupulin); l/irlZ
28/9 (RPSC). BOLIVIA . Consata. C. Luer S. 1I .
(SEL, Ihree speci mens). Santa Cruz: Chiquitos,
Robaré. VlIsqllez 895 (Herb. Vasquezianum): Tu mupasa, White / /20 (A MES); Rurrcnabaque,
Wllite 2323 (AMES). PER U. Mono Calzada, D.E.
Be/lllett 3245 (SEL). Without collcction dala:
Hor/. Linden (K); cIIII. Hort. Bot. KelV (K); Han.
While ( K ); l/orr. Samler (K); l/orl. Ma/llin (K);
Hort. Filler ( K ).
4
4
T. pallduralum Schweinf.. Amer. Orcl/id SOl.'.
Bu/l. 14: 104. 1946. TYPE: PER U. San Manín:
Juan Jui, Alto Río Huall¡¡ga, C. Klug 4260 (Ho101ype. AMES: lsolype. K).
DlSTR IB UTION : Endemic 10 Peru .
REP RE SENTATIVE S PECIM ENS: PERU :
San Manfn: Huallaga, Bellavista, J. Sc/lIIl/ke V.
sub D. Bennelt 1/35: Moyobamba. R. Stiimpjfe
s. /!. (pholo in Teuscher, 1964).
T. purpllreum Lindl. ex Rehb.r., C(ml. C/¡ron .
772. 1854. TYPE: GUYANA. Ex ¡lOrt. Veitch
(HoloIYpe, K).
DI STRIB UTION: Endemic 10 Guyana.
T. r ecunum Lindl. , EdlVanJs's Ba r. Reg. 29:
misc. 9. 1843. TYPE: GUYANA. Ex horr. Loddiges S. I/ . (Holotypc, K!).
203
T. comU41'(l(,:cae Rchb.r. in Reichen bac h
Herb. , M ....
T. pleclrophora Rchb.f. in Herb. Jard . Bol.
Etal Bruxelles ( BR), M.f.• l/ O /! (Lodd. ex Undl. )
Rehb.f. ["" Pleclrop!tora iridifolial.
DISTRIB UTION : Ecuador (7), Guyana and Su+
rinam.
REPRESENTATIVE S PEC IMENS : ECUADOR. Moro na 4Sant iago : Mi s ión Bomboi za,
f/olm-Niel.'iell el al. 4/97 (nol seen, but very dubious). SURINAM. Paramaribo, 27 Feb. 1858,
Focke S./I. (W). Concordia : Saramacca, without
collector (as T. conlll-vaccae, BR); same locality,
without collector (BR). Without locali ly. lVl/Useh laegel .... II. (W); cult. DIlII .~.II. (W).
T. tenuiflorum Undl.. Paxr. FlolV. Carll. 1: 12.
1850. TYPE: BRAZIL Bahia: Wi thout locality.
M orel s.n. (Holotype, K).
DlSTRIBUTION: Brazil.
TricllOCetltrllm wagneri Pupulin sp. I/OV. TYPE:
BRAZIL Wilhout locality, inlroduced by A. Seidel. flowered in cuttivation by A. Wagner in Sanla
Margherita, Italy. 25 Sept. 1990. F. PII/Jlllin 289
(Hololypc, SEL). Fig. 14.
Species ad Triehocenlro fu sco Undt. simili s.
sed calcan mul lo magis longiore quam labclli. labello oblongo-panduralO rotundato marginibus
lev iter crenulati s, calli s parallelis et alis columnae
brunneo slriat is dirrert.
Plant epiphytic. ce... pitose, pendent 10 SUbcl\.'1:l,
wi th abbreviatcd rhizome. Rools filironn, fl exuous, glabrous, silvery whi te with grccn apex. Pseu4
dobulbs subglobose. up 10 5 mm long, 3.5-4.5
mm wide, mOllophyllous. Leaf ovate 40blong to cl 4
liptic<>()blong, retuse. narrowed to a conduplicate.
sessile base. light grecn, 6-1 1 cm long, 3-3.5 cm
wide. lnflol'cscence lateral. basal. pendem, successive, I- many-Howercd: peduncle terete. 7- 9 cm
long, concealed by 2-3 eoncave, ovate, papir¡¡ceous bracts, 10 8 mm long. Ovaries linear4c1avate,
tercte, abaut 4 cm long including Ihe pedicel.
flowers with rree scpals and petals, du ll brown
with yellowish apexes and ivory while lip, m¡¡rked
near the b¡¡se by yellow with purple-brown radiating lines and blotches. Dorsal sepal oblanceolate. acule. concave toward the apex, to 20 mm
long, 6.5 mm wide. Lateral sepals obl iquely subfalcate, acule 10 subobluSC, subcari nale low¡¡rds
the apex and slightly concave, 10 2 [ mm long, 5-
PUPULlN- A REVISION OF TR1CHOCEIVI"RUM
5 mm
1 cm
Fig. 14.
5 ,11m
Trichocenlrum ....agneri Pupulin. Voucher: Wilholll CQll ect ion data. introduccd by A. Scidcl. flowcred in cu lli val ion by
A. Wagner. F. Pupulin 289 (SEL).
204
PUPULIN- A REV IStON OF TR/CHOCI:.'NTRUM
5.5 mm wide. Pela ls linear-obovate. obtuse. sorneli mes slightly concave towards Ihe apex. 18.5 mm
long, 5.5 mm wide. Lip from a cuneate base oblong-subpandurate, adnale to the column . to 26
mm long. 11 .5 mm wide. emarginale in front. wi th
plane 10 slighlly crenu late margins. producing with
the column base a sle nder. linear-attenualc. fexuous
spur. 5 cm long. 0.35 cm wide ¡II the base; disc
with four slender. puberu lenl keels. the exlernal
ones shorter. extending parallcl from the base to
the middle of the lip. Column short, Slout. withoul
a foot. 10 6 mm long, with a pair of largc. crecl,
c uneate. s ubquadrate. irregularly erose-dentate
wings. completely striped with brown . Anlher
whitc. subglobosc. glabre. Pollinia 2, pyriform.
deeply concave. on a short obtriangu lar sti pe; viscidium pel lale. brown.
ETYMOLOGY: Named in honor of Alessandro
Wagner. of Mila n. Italy, who firsl fl owered the
species in his colleclion,
D1 STRIBUTION: Brazil.
REPRESENTATIVE SPEC IMENS: BRAZIL.
Withoul colleclion dala. inlroduced by A. Sei del
and flowc red by F. Pupulill 290 (Herb. Pupu li n);
cu ltivaled by A. Riboni, F. PI/pulin 29/ (Herb.
Pupulin).
The species was introduced in lIal y in early
summcr 1989 IOgcther with a group of T. fuseum
plants com ing from Brazi l. Several specimens
Howered in cultivalion in 1990. showing flowers
thal onl y superficially rescmble those of T. lu.~­
eum Lindl. Tricl/Oeel/lmm IVtlgl/eri may be easily
dislinguished from T. fuseul/I mainly on Ihe basi s
of spur length Wh~'is twice longer than Ihe !ip,
whcreas it is as 1 g as the lip or slight ly shorter
in T. I II.I'Cllm. The ip is oblong. plane, only slightIy crcnulale lowards e middle. It presents on the
disc four slender. puberu lenl keels. thal mn parallel from the e ntrance of the spur to the midd le
of Ihe lamina. Column wings lIre wide. cunellle,
subqadrate and concave; Ihey are intensively
striped by brown, whereas they are slighlly Spolted in T./lIs(."l/l/I. The species fl owers in lale autumn , fro m Septcmbcr through Decembcr.
•
i
"".
l..
. .~
'y.
'.
.." ...
-
'\
i.,
",.,
....
,,".'
!
. 1. b'rachyi;Qras
• l . brCtViealcara!Um
[,ig. 15. Dislribulion of Ihe spccies of Trichocemrum bru ch)"ceras group.
cll)'ce ra.~ fl owers. as we ll as these of T. brevicalc:aratum C. Schweinf., presenl a narrow, deeply
concavc lip wi th ac utc apex and short, tubu lar
spur. Both the high ly rcvolule margins of the \ip
and ils acule lo acuminate apex are useful c haracte rs 10 di slingui sh the taxa of Ihis group from
Ihe main body of the genus. The two species can
be easily distingui shed by Ihe length of Ihe lip: ir
is longer Ihan the sepals and wi th Iwo keels near
¡he base in T. brachyc:eras. while it is marked ly
shorter than the sepals and wi thoul keel s in T.
brevicalclIratum. NOIhing is known about the
vegelative morphology of T. bracllyceras. bUI T.
brevicalcaratum prescnts a Iypical habit for Ihe
genus, with short, I -fl owered inflorcscences.
Tricl,ocefllrllm bracllJlceras group
Key lo Ihe species or
Trichocenlrum brachyceras gl'oup
Only two species belong in Ihis group, one e ndemic 10 Colombia and the other 10 Pem (Fig.
15). Tric:lwcellTml/l brachyeeras Schltr. is known
only from a Schlcchler's tracing of Ihe Iype (pu blished by Mansfeld. 1929). Tricllocelllrum bra-
1u. Lip longc r lhan scpals. apiculale. wilh IWO disli nCI
keels !lear lhe base
. , . , 1. brach)'ceros
lb. Lip mar1.:edly shorter lhan sepa!s. minulcly reluse.
Wi lhoul kecls
. ... .. T. brevicalcarO/um
205
PUPULlN- A REVIS ION OF TRICHOCE.NTRUM
Fig, 16.
Trichocrmnm, brr"icolcort'WIII Schweinf, PERU. Junín: Chaochamayo Va lley. C. Sch,,,,l:;r :1.11. (from Schweinrurth.
1945),
T. bracllyceras 5chllr.. Reperr. S/Jec. Nov. Regni
Veg. Bei/¡. 7: 184. 1920. TYPE: COLOMBIA.
Cauca: M. Madero .\'./1. (Holotypc. B. destroyed;
drawing. Reperr. Spe(;. Nov. Regui Veg. Bei/I. 57:
t. 65. Fig. 252. 1929).
DISTRIBUTION: Endemic 10 Colombia.
Chanchamayo Valley. 1.800 m. July " 19241921". C. Scllllnke s.n. (Holotypc. F).
DISTR IBUTlON: Endemic (O Peru.
REPRESENTATl VE SPECIMEN5: PER U. Junín : Chanchamayo Val ley. C. S(:llIlIIke 565 (F).
Trichocelltrum tigrillum group
Trichocelllrum tigrinum. a species from Ecuadorean and nonhem Peruvian Paci fi c regions is
T. brevicalcaralum Schweinf.. Amer. Orc:hid Soco
Bull. 13: 388. 1945. Fig. 16. TYPE: PERU. Junin.
206
PUPULlN- A REVISION OF TRICHOCI::NTRUM
(SEL); e. H. Dad.wII & LB. Thiel/ /744 (SEL).
Prov. Guayas: Manglaralto. C H. Dodson 16252
(RPSC); Cordi llera de Colonehe-Chongón. 1500
m, E. Sanrie ...leball sub F. PU/JIllin 287 (Herb.
Pupulin). El Oro: Santa Rosa. E.F. A/ldré 4.17/
(K): F.e. Lehmann /000 and /001 (K); Río Calara. bctwcen Ayabám and Zaruma. F.e Lehmallll VI. (K). Prov. of Loja: Catamayo (K); Macará. P. Andreel/a .".n. (Herb. Pupulin). Without
coJlection data. ex Hort. Lemujle (He rb. Pupulin).
PERU. Piura: Ayabaea. Yanehal a. D. Bemleft e/
al. 34 /5. Wi lhout eollection data. ex Hart. Ribolli
S.II. (Herb. Pupu li n).
L.
j
"
\
EXC LUOEO SIIECI ES
•
FJg. J?
T. (llalllm Rolfe. Bull. Müc. Inform. 197. 1890.
TYPE: COLOMB IA. Mil/iean .1". 11. :::: Pleelrophora alala (Ro lfe) Garay. BOl. Mu.". LeaJI. 21: 261,
1967.
T. fUI/ale (Swartz) Lindl. , Edwards's BOl. Reg .
23: sub 1. 1951. 1837 . TYPE: JAMA ICA. Swartz
s.n. (Hololypc of Epidendrll111 fimale. BM ) =
Dendrophylax funali ... (Swartz) Fawc .. Fl. PI.
l amo 40, 1893.
T. iridifolillm Lodd. ex Lindl. . Edwan/.f·.\· Bot.
Reg. 24: Mise. 94, 1838. TYPE: SURI NAM.
Wull...clrfaege/ /805 = Pleelrop/¡ora iridifolia
(Lodd. ex Lindl.) Focke. Tljdschr. Wis.'''. en Naturk. AnlSlerd. 1: 212. 1848.
T. plectro/J/rora (Lodd. ex Lindl.) Rehb.f.. Anll.
Bol. Sysl. 6: 544. 1863. = Pleclrop/¡ora iridifolia
(Lodd. ex Lindl.) Focke. Tijd.fc/¡r. Wiss. ell NaIIlrk. Afm.·lerd. 1: 212. 1848.
T. Iriqlle/mm Rolfe. Gard. Chron. ser. 3. 9:
70 1, 1891. TYPE: PERU. Ex Hort. Clwrle,l'Ivvrrli
= Plec:tro/Jlrora Iriquetra (Rolfe) Cogn., FI. Bras.
3: parto 6: 184.1904.
••
•
OislribUlion of Triclwcenlrllln lig, ¡n"m.
the SQle spccies referred 10 Ihis group (Fig. 17).
The very spreading flowers of this species, (he
showing coloration and sweel seent, which descrvcd il Reichenbach's recognilion as " Trichocentrurn princeps." can easily distinguish th is
11l0nolypic group. According lo van der Pijl and
Dodson ( 1966), T. ligrillllm is poll inaled by Eulaema cin811lata. The modified. very thick leaves
of T. /igr;,wm a llow the plants of this specics 10
mantain a hydrie balance in the sub-arid scasonal
foresls of the coastal plains of EcuadOr and Peru ,
whcre Ihe dry season may execcd five months.
CHEC KLlST OF SPECIES
ANo SYNONYMS
Acoidium filSCllf1l Lindl . = T. fi,selml Lindl.
Ore/lis pUllcta/a Ruiz & Pavón = T. IJlde/lrum
Pocpp. & Endl.
T. alalum Rolfe = Plectro/J/lOra (1 /ata (Ro lfe)
Garay
T. albif/orum Ro lfe = T. candidllm Lindl.
T. albQ-coccineum Linden
T. albo-¡Jllr/JIlreum Linden & Re hb.f. :: T. albocoecineum Linden
T. albo-purpurewlI varo Slrialllm Linden & Roo.
= T. albo-co(:cineum Linden
T. ligrinwn Linden & Rehb.f.. Gard. Chroll. 862.
1869. TYPE: ECUADO R. Guayas: G uayaquil,
1875. Wa//ü S.II. (Ho lo lypc. W). Fig. 18.
r. ligrillum varo ~;ple l"Jell .\· Linden & Rod ..
Un(/enia 1: 53, L 24. 1885.
DISTR1BUTION : Ecuador and Pcru.
REPRESENTATIVE SPEC IM ENS: ECUADOR. Manabí: MonticriSli, cast of Manta. e H.
Dods()II / /3 and 340 (SEL). e H. Dodsoll s.n.
207
PUPULl N- A REVISION OF TRICHOCENTRUM
5mm
2cm
3cm
5mm
3cm
Fig. 18.
Trichocen/rum /igrinum Lillde ll & Rchb.f. VOl.lcher: ECUADOR. Loja: Macará. I~ Atldree/w
208
VI.
( Herb. Pupulill).
PUPULl N-A REVI SION OF TRICHOCENTR UM
T. alboviolaceum Sch lu. = T. albo-coccineum
Linden
T. amawnicum Barb. Rodr. = T. fuscum Lindl.
T. brachyceras Schltr.
T. brandtiae Krzl.
T. brenes;; Schllr.
T. brevicalcaratum Schweinf.
T. caloceras Endres & Rchb. f.
T. candidum Lindl.
T. capistratum Linden & Rchb. r.
T. cornucop;ae Li nden & Rchb.f. = T. fu.~cum
Lindl.
T. cornu-vaccae Rchb.f. = T. recurvum Lindl.
T. costaricense Mora-Retana & Pupulin
T. cymbiglossum Pupulin
T. dianthum Pupulin & Mora-Retana
T. estrellense Pupulin & J.B. Gurda
T. funale Lindl. = Dendropllylax funa lü
T. fu scum Lindl.
T. har/ii Rolfe = T. f uscum Lindl.
T. hoegei Rchb,f.
T. ;onophtalmum Rchb.f. = T. albo·coccineum
Linden
T. iridifo/ium Lodd ex Lindl. = P/ectrophora irid/folia (Lodd. ex Lindl. ) Pode
T. leeanum Rchb.f. = T. albo-cocc;inewn Linden
T. longicalcaratum Rchb.f.
T. macula/um Lindl . = T. pulcllrum Poepp. &
Endl.
T. mattogrosserue Hoehne = T. fuscum Lindl.
T. orthoplectron Rchb.f. = T. albo-coccineum
Linden
T. panamense Rolfe = T. capislratllm Linden &
Rchb.f.
T. panduratum Schweinf.
T. pfavii Rchb.f.
T. pfav;; varo zonale Rchb.f. = T. pfav;; Rchb.f.
T. f)fav;; f. album Henderson = T. pfavi; Rchb.f.
T. pineU Lindl. = T. fuscu m Lindl.
T. plectrop/lOra (Lodd. ex Lindl. ) Rchb.f. = P/ectrophora iridifoUa (Lodd. ex Lindl.) Focke
T. plectrop/IOTa Rchb.f. = T. recurvum Lindl.
T. pOTf)hyrio Rchb.f. = T. albo-coccineum Linden
T. pulchrum Poepp. & Endl.
T. purpureum Lindl. ex Rchb.f.
T. pu.~i//um Lehm . = T. capislratum Linden &
Rchb.f.
T. recurvum Lindl.
T. pfavii
T. saundersianum Endres & Rchb.f.
Rchb.f.
209
T. ,mundersii Endres & Rchb.f. = T. pfavii
Rchb,f.
T. speciosum Schlim = T. pu/chrum Poepp. &
Endl.
T. tenuiflorum Lindl.
T. tigrinum Rchb,f.
T. tigrinum varo splendens Linden & Rod. = T.
ligrinum Rchb,f.
T. trique/rum Rolfe = PlectropllOra triquetra
(Rolfe) Cagn.
T. verruciferum Schhr, = T, brandliae Krzl.
T. wagneri Pupulin
T. zonale Rchb.f. = T. pfavii Rchb.f.
LITERA TURE C IT ED
Beáner. J. 1963. 111e gcnus Trich(Jct'nlrulII. Amer. Orchid
Soc. Bull. 32: 194- 1%.
Bentham. G. 1881. Notes on Orchideae. J. Uf"'. Soc .. BOl.
18: 281-360.
Bracm G. J. 1993. Studies in the Oncidiinac. Discussion of
sorne taxonomic problcms with description of Gudrunio
Braem, sell. '101'.• and reinstatement of the genus Lop/¡io f'
is Rafi nesq ue. ShlechreriOllO 4: 8-2 1.
Cabello Martfn, M .. M. L. Garc(a·Oclloo Roldán, alld P. Nájera
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al VirreinolO de! Perú (1777- 1788), Lunwerg and Real
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Cllase, M. W. 1986. A reapprdisal o( the Ollcidioid orchids.
S)'SI. Bol. 11: 477--491 .
--o 1986b. A monograph of Lel)(:hilus (Orchidaceae). 5)'51.
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Chasco M. W.. and J. D. Palmer. 1992. Floralmorphology and
chromosome number in subuibe Oncidiinae (Orch idaceae): evolutionary insights frorn a phylogenetic analysis
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síficalio'J, Harvard University Press, Cambridge, Massa·
chusetts.
--o 1993. Phylogeny (lIJ(J classíficatiolJ r>/lhe orchidlamily.
Dioscoridcs Press. PonlarKi.
Dresslcr. R. L .. and C H. DodSOIl. 1960. CJassilkalion ;md
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e¡ue!(J. AII iIIJ/Slraled field guide. Botanical Museum,
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KrJnzlill, F.W.L. J 897. Orchid(Jceae nO\'{le. Bul/. Herb. 8oiss.
5: 109.
Lindley, J. 1843. Trich(JCf!nlrUIII recul"i'J/m. Edwards's 801.
Reg. 29: Mise. 9.
1850, Trichoct:mrum lenuij/orllm. Paxton's FI(1w.
Gard. 1: ]2.
--o 1854. Trichocelllrum pille!i. G(Jrd. Chron. 772.
Poeppig, E. F., and S. L. Endl icher. J 836, Trichoce'JlfI"" IJUI·
chrum. No~. Gell. Sp. PI. 2: 11. t. 115.
PUPULlN-A REVIS ION OF TRfCIIOCENTRUM
Ro lfe, R. A .. 1893. Trichocenfrwn olbiftorum. 8/111. Mise. In ·
fonn. 337.
1894. Trichoctnfrum hanli. Bu/l. Mise. flr/onu . 395.
--o 1913. Triehoce/JIrmn panomense. 8,,11. Mise. Inform.
341.
Schlcc hler. R. 1923. 7"richocenm,m brenesll. NtfH' rl. SfJtC.
No~. Negni Velo'. IJeih. 19: 248.
--o 1926. Das Syslcm der Orchidacecn. Nurizj}/. /JOI. Gan
Mus. Berlin- f)lIhlem 9: 563-591.
Schwcinfurth. C. 1945. A remarkabl c orchid fmm ¡he Andcan
region. Amer. Orchid Soc. 8ull. 13: 388.
--o 1946. A novelty in Peruvian Triehocen""m. Alllu. Oro
chid Soc. 8u/l. 14: 1M.
--o 1959. Key lo lhe on: hids. in Wilhncr C.L. (ed.), TIre
orchids. A scie/JIijie sun't)'. J. Wiley ami Sonso New York.
Te uscher, H. 196 !. Some Trichocentro ms. Amu. Orchid SiK.
8ull. 30: 389.
--o 1964. 7"rlchocentrum IXJndur(llum. Alller. Orchitl Soc.
/Jull. 33: 644-646.
van der Pijl. L. and C. H. Dodson. 1966. On:/riIJ F/a ....ers.
Tlreir Pollilllj/imr (l/JI! E.·olrtlivn. Universi l)' of Mi;uni
I'ress. Coral Gabl es.
Pu pul in, F. 1993. Note sul genere Trichoc~nln4m. l. Tricho·
C~nfn4m caplslrolllm Linden & RchbJ. Orch. Sponf. Col/.
6: 27- 30.
--o 1993b. Note sul genere Trichoct'rll rum. 2. Trichoc~n ­
"rllllfiucum Lindl.: una specie nuova per la flora dell" Ecuador. Orch. Sfxml. Cok 6: 53-59.
1994. A ne w spedes of Tr!chrKtmlrrmr fmm Costa
Rica. Umllt)"mra 9: 51-53.
--o and 1). E. Mora- Retana. 1994. A rev ision of lhe Cosla
Rican Trichocenlrum species (On:hidaccae). Selb)"ana 15:
87- 103.
Rcichenbaeh, H. G. 1869. Trichocenln4m romrrcoplae. Rtf
&t. 2: sub l. 77.
--o 1876. Triehocen/rum lonoph/almum. Gard. Chron. 100.
188 !. Trichocenlrum hcwgei. Gard. Chron. n. s. 16:
7 17.
1883. Trichoc~nfn4m onhopleclrun. Gllnl. Chron. n. s.
19: 562.
Rodrfguez Caballero. R. L. 1986. Gbum1s de IIrquMells de
Cosw Nica. San José. Editorial Uni vcn; idad de Cosla
Rica.
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